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Ribeiro, V., Mateus O., Holwerda F., Araújo R., & Castanhinha R. (2014).  Two new theropod egg sites from the Late Jurassic Lourinhã Formation, Portugal. Historical Biology. 26(2), 206-217. Abstractribeiro_et_al_2014_theropod_eggs_nest_portugal.pdfWebsite

Two new Late Jurassic (uppermost Late Kimmeridgian) dinosaur eggshell sites are described, Casal da Rola and Porto das Barcas, both near Lourinha˜, central-west Portugal. Casal da Rola yields eggshells with an obliquiprismatic morphotype comparable to those from a nest with the associated fossil embryos from Paimogo, tentatively assigned to the theropod Lourinhanosaurus antunesi. The Porto das Barcas eggshells have a dendrospherulitic morphotype with a prolatocanaliculate pore system. This morphotype was also recognised in eggshells from a clutch with associated Torvosaurus embryos at the Porto das Barcas locality. A preliminary cladistic analysis of eggshell morphology suggests theropod affinities for the Casal da Rola eggs, but is unable to resolve the phylogenetic position of the Porto das Barcas eggs. The eggshells at both sites are preserved in distal flood plain mudstones and siltstones. Carbonate concretions within the deposits indicate paleosol development.

Ribeiro, V., Mateus O., Holwerda F., Araújo R., & Castanhinha R. (2014).  Two new theropod egg sites from the Late Jurassic Lourinhã Formation, Portugal. Historical Biology. 26, 206-217., Number 2 Abstract
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Ribeiro, V., Holwerda F., & Mateus O. (2013).  Theropod egg sites from the Lourinhã Formation, Portugal. Journal of Vertebrate Paleontology, Program and Abstracts, 2013. 198.ribeiro_et_al_2013_theropod_egg_sites_lourinha_svp_meeting_abstracts_213.pdf
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Holwerda, F., Tschopp E., & Mateus O. (2014).  Sauropod body fossils in Europe: overview and current issues. XII EAVP Meeting XII Annual Meeting of the European Association of Vertebrate Palaeontologists – Abstract Book. p.77., Torino 24-28 June 2014holwerda_et_al_2014_sauropods_europe_eavp.pdf
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Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ignacio-Ruiz Omeñaca J., Lepage Y., Havilk P., & Andrade M. B. (2014).  Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia). Royal Society Open Science. 1(140222), 1-42.young_et_al_2014_machimosaurus_crocodylomorph_revision.pdf
Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ruiz-Omeñaca J. I., Havlik P., Lepage Y., & De Andrade M. B. (2014).  Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia). Royal Society Open Science. 1, , Number 2 Abstract
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Hendrickx, C., Mateus O., & Araújo R. (2015).  A proposed terminology of theropod teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. e982797. Abstracthendrickx_et_al_2015_theropod_teeth_svp.pdfWebsite

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Hendrickx, C., Mateus O., & Araujo R. (2015).  A proposed terminology of theropod teeth (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 35, , Number 5 Abstract
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Ceríaco, L. M. P., Gutiérrez E. E., Dubois A., Abdala C. S., Alqarni A. S., Adler K., et al. (2016).  Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences. Zootaxa. 4196(3), 435 - 445., 2016 AbstractWebsite
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Ceríaco, L. M. P., Gutiérrez E. E., Dubois A., Abdala C. S., Alqarni A. S., Adler K., et al. (2016).  Photography-based taxonomy is inadequate, unnecessary, and potentially harmful for biological sciences. Zootaxa. 4196, 435-445., Number 3 Abstract
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Hendrickx, C., Hartman S. A., & Mateus O. (2015).  An overview of non-avian theropod discoveries and classification. PalArch’s Journal of Vertebrate Palaeontology. 12(1), 1-73. Abstracthendrickx_etal_2015_non_avian_theropods_pjvp12_11.pdfWebsite

Theropods form a taxonomically and morphologically diverse group of dinosaurs that include extant birds. Inferred relationships between theropod clades are complex and have changed dramatically over the past thirty years with the emergence of cladistic techniques. Here, we present a brief historical perspective of theropod discoveries and classification, as well as an overview on the current systematics of non-avian theropods. The first scientifically recorded theropod remains dating back to the 17th and 18th centuries come from the Middle Jurassic of Oxfordshire and most likely belong to the megalosaurid Megalosaurus. The latter was the first theropod genus to be named in 1824, and subsequent theropod material found before 1850 can all be referred to megalosauroids. In the fifty years from 1856 to 1906, theropod remains were reported from all continents but Antarctica. The clade Theropoda was erected by Othniel Charles Marsh in 1881, and in its current usage corresponds to an intricate ladder-like organization of ‘family’ to ‘superfamily’ level clades. The earliest definitive theropods come from the Carnian of Argentina, and coelophysoids form the first significant theropod radiation from the Late Triassic to their extinction in the Early Jurassic. Most subsequent theropod clades such as ceratosaurs, allosauroids, tyrannosauroids, ornithomimosaurs, therizinosaurs, oviraptorosaurs, dromaeosaurids, and troodontids persisted until the end of the Cretaceous, though the megalosauroid clade did not extend into the Maastrichtian. Current debates are focused on the monophyly of deinonychosaurs, the position of dilophosaurids within coelophysoids, and megaraptorans among neovenatorids. Some recent analyses have suggested a placement of dilophosaurids outside Coelophysoidea, Megaraptora within Tyrannosauroidea, and a paraphyletic Deinonychosauria with troodontids placed more closely to avialans than dromaeosaurids.

Hendrickx, C., Hartman S. A., & Mateus O. (2015).  An overview of non-avian theropod discoveries and classification. PalArch’s Journal of Vertebrate Palaeontology. 12, 1-73. AbstractWebsite

Theropods form a taxonomically and morphologically diverse group of dinosaurs that include extant birds. Inferred relationships between theropod clades are complex and have changed dramatically over the past thirty years with the emergence of cladistic techniques. Here, we present a brief historical perspective of theropod discoveries and classification, as well as an overview on the current systematics of non-avian theropods. The first scientifically recorded theropod remains dating back to the 17th and 18th centuries come from the Middle Jurassic of Oxfordshire and most likely belong to the megalosaurid Megalosaurus. The latter was the first theropod genus to be named in 1824, and subsequent theropod material found before 1850 can all be referred to megalosauroids. In the fifty years from 1856 to 1906, theropod remains were reported from all continents but Antarctica. The clade Theropoda was erected by Othniel Charles Marsh in 1881, and in its current usage corresponds to an intricate ladder-like organization of ‘family’ to ‘superfamily’ level clades. The earliest definitive theropods come from the Carnian of Argentina, and coelophysoids form the first significant theropod radiation from the Late Triassic to their extinction in the Early Jurassic. Most subsequent theropod clades such as ceratosaurs, allosauroids, tyrannosauroids, ornithomimosaurs, therizinosaurs, oviraptorosaurs, dromaeosaurids, and troodontids persisted until the end of the Cretaceous, though the megalosauroid clade did not extend into the Maastrichtian. Current debates are focused on the monophyly of deinonychosaurs, the position of dilophosaurids within coelophysoids, and megaraptorans among neovenatorids. Some recent analyses have suggested a placement of dilophosaurids outside Coelophysoidea, Megaraptora within Tyrannosauroidea, and a paraphyletic Deinonychosauria with troodontids placed more closely to avialans than dromaeosaurids.

Hendrickx, C., Hartman S. A., & Mateus O. (2015).  An overview of non-avian theropod discoveries and classification. PalArch{'}s Journal of Vertebrate Palaeontology. 12, 1-73., Number 1 Abstract
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Hendrickx, C., Hartman S. A., & Mateus O. \á\}vio (2015).  An overview of non-avian theropod discoveries and classification. PalArch\’\}s Journal of Vertebrate Palaeontology. 12, 1-73. Abstract
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Hendrickx, C., Hartman S. A., & Mateus O. \á\}vio (2015).  An overview of non-avian theropod discoveries and classification. PalArch\’\}s Journal of Vertebrate Palaeontology. 12, 1-73. Abstract
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Hendrickx, C., & Mateus O. (2012).  Ontogenetical changes in the quadrate of basal tetanurans.. 10 th Annual Meeting of the European Association of Vertebrate Paleontologist ¡Fundamental! . 20, 101-104. Abstracthendrickx__mateus_2012_ontogenetical_changes_in_the_quadrate.pdf

Although nonavian theropod have received considerable interest in the last years, their ontogeny still remains poorly understood, especially the ontogenetical changes affecting their skull (Rauhut
and Fechner, 2005). The quadrate, for instance, is preserved in several embryos and juvenile specimens belonging to many clades of theropods such as the Tyrannosauridae (Carr, 1999), Compsognathidae (Dal Sasso and Maganuco, 2011), Therizinosauroidea (Kúndrat et al., 2007), Oviraptoridae (Norell et al., 1994; Norell et al., 2001; Weishampel et al., 2008) and Troodontidae (Varrichio et al., 2002) but very little is usually said about the anatomy of this bone and no one has ever investigated ontogenetical variation in the nonavian theropod quadrate. The discovery of two quadrates belonging to embryos of the sinraptorid Lourinhanosaurus antunesi from Portugal and five isolated quadrates pertaining to juvenile, subadult and adult specimens of Spinosauridae from Morocco fills this gap and allows some ontogenetic information to be drawn for this bone in these two specific clades of Theropoda.
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Hendrickx, C., Araújo R., & Mateus O. (2012).  The nonavian theropod quadrate: systematics usefulness, major trends and phylogenetic morphometrics analysis. : Journal of Vertebrate Paleontology, Program and Abstracts, 2012, p.110. ISSN 1937-2809 Abstracthendrickx_araujo__mateus_2012_quadrate_theropods_svp_2012_abstract_book_meeting_abstracts.pdf.pdf

The quadrate in nonavian theropods is incredibly diverse morphologically; however this morphological disparity has been underestimated for taxonomic purposes. The quadrate topological homologies and anatomy, as well as the terminology, among nonavian theropod clades are reviewed. In order to evaluate the phylogenetic potential and investigate the evolutionary transformations of the quadrate, we conducted a Catalano-Goloboff phylogenetic morphometric analysis using 3 morphometric characters, a total of 28 landmarks coded for 23 taxa, as well as a cladistic analysis using 115 discrete quadrate-related characters coded for 43 taxa. The cladistic analysis provides a fully resolved tree mirroring the current classification of nonavian theropods. The quadrate morphology by its own provides a wealth of data with strong phylogenetic signal. Several unambiguous synapomorphies support nonavian theropod relationships and the resulting consensus tree allows inference of major trends in the evolution of this bone. Important synapomorphies include: for Abelisauridae, a lateral ramus extending to the ectocondyle; for Tetanurae, the absence of the lateral process; for Spinosauridae, a medial curvature of the ventral part of the pterygoid ramus occurring just above the mandibular articulation; for Neotetanurae, an anterior margin of the pterygoid flange formed by a roughly parabolic margin; and for Tyrannosauroidea, a semi-oval pterygoid flange shape in medial view. The Catalano-Goloboff phylogenetic morphometric analysis reveals two main morphotypes of the mandibular articulation of the quadrate linked to function. The first morphotype, characterized by an anteroposteriorly broad mandibular articulation with two ovoid/subcircular condyles roughly subequal in size, is found in Ceratosauria, Tyrannosauroidea and Oviraptorosauria. This morphotype allows a very weak displacement of the mandible laterally. The second morphotype is characterized by an elongate and anteroposteriorly narrow mandibular articulation and a long and parabolic/sigmoid ectocondyle. Present in Megalosauroidea, Allosauroidea and Dromaeosauridae, this morphotype permits the lower jaw rami to be displaced laterally when the mouth opened.

Hendrickx, C., Araújo R., & Mateus O. (2014).  The nonavian theropod quadrate II: systematic usefulness, major trends and cladistic and phylogenetic morphometrics analyses. PeerJ PrePrints. 2, e380v2., 2014 AbstractWebsite

The skull-bone quadrate in nonavian theropods is very diverse morphologically alongside the disparity of the group as a whole. However this disparity has been underestimated for taxonomic purposes. In order to evaluate the phylogenetic potential and investigate the evolutionary transformations of the quadrate, we conducted a Catalano-Goloboff phylogenetic morphometric analysis as well as a cladistic analysis using 98 discrete quadrate related characters. The cladistic analysis provides a fully resolved tree mirroring to some degree the classification of nonavian theropods. The quadrate morphology by its own provides a wealth of data with strong phylogenetic signal and allows inference of major trends in the evolution of this bone. Important synapomorphies include: for Abelisauroidea, a lateral ramus extending to the ectocondyle; for Tetanurae, the absence of the lateral process; for Spinosauridae, a medial curvature of the ventral part of the pterygoid ramus occurring just above the mandibular articulation; for Avetheropoda, an anterior margin of the pterygoid flange formed by a roughly parabolic margin; and for Tyrannosauroidea, a semi-oval pterygoid flange shape in medial view. The Catalano-Goloboff phylogenetic morphometric analysis reveals two main morphotypes of the mandibular articulation of the quadrate linked to function. The first morphotype, characterized by an anteroposteriorly broad mandibular articulation with two ovoid/subcircular condyles roughly subequal in size, is found in Ceratosauria, Tyrannosauroidea and Oviraptorosauria. This morphotype allows a very weak displacement of the mandible laterally. The second morphotype is characterized by an elongate and anteroposteriorly narrow mandibular articulation and a long and parabolic/sigmoid ectocondyle. Present in Megalosauroidea, Carcharodontosauridae and Dromaeosauridae, this morphotype permits the lower jaw rami to be displaced laterally when the mouth opened.

Hendrickx, C., Aráujo R., & Mateus O. (2015).  The non-avian theropod quadrate I: Standardized terminology with an overview of the anatomy and function. PeerJ. 2015, , Number 9 Abstract
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Hendrickx, C., Bell P. R., Pittman M., Milner A. R. C., Cuesta E., O'Connor J., Loewen M., Currie P. J., Mateus O., Kaye T. G., & Delcourt R. (2022).  Morphology and distribution of scales, dermal ossifications, and other non-feather integumentary structures in non-avialan theropod dinosaurs. Biological Reviews. , Number n/a Abstracthendrickxetal.2021.morphologyanddistributionofscales.pdfWebsite

ABSTRACT Modern birds are typified by the presence of feathers, complex evolutionary innovations that were already widespread in the group of theropod dinosaurs (Maniraptoriformes) that include crown Aves. Squamous or scaly reptilian-like skin is, however, considered the plesiomorphic condition for theropods and dinosaurs more broadly. Here, we review the morphology and distribution of non-feathered integumentary structures in non-avialan theropods, covering squamous skin and naked skin as well as dermal ossifications. The integumentary record of non-averostran theropods is limited to tracks, which ubiquitously show a covering of tiny reticulate scales on the plantar surface of the pes. This is consistent also with younger averostran body fossils, which confirm an arthral arrangement of the digital pads. Among averostrans, squamous skin is confirmed in Ceratosauria (Carnotaurus), Allosauroidea (Allosaurus, Concavenator, Lourinhanosaurus), Compsognathidae (Juravenator), and Tyrannosauroidea (Santanaraptor, Albertosaurus, Daspletosaurus, Gorgosaurus, Tarbosaurus, Tyrannosaurus), whereas dermal ossifications consisting of sagittate and mosaic osteoderms are restricted to Ceratosaurus. Naked, non-scale bearing skin is found in the contentious tetanuran Sciurumimus, ornithomimosaurians (Ornithomimus) and possibly tyrannosauroids (Santanaraptor), and also on the patagia of scansoriopterygids (Ambopteryx, Yi). Scales are surprisingly conservative among non-avialan theropods compared to some dinosaurian groups (e.g. hadrosaurids); however, the limited preservation of tegument on most specimens hinders further interrogation. Scale patterns vary among and/or within body regions in Carnotaurus, Concavenator and Juravenator, and include polarised, snake-like ventral scales on the tail of the latter two genera. Unusual but more uniformly distributed patterning also occurs in Tyrannosaurus, whereas feature scales are present only in Albertosaurus and Carnotaurus. Few theropods currently show compelling evidence for the co-occurrence of scales and feathers (e.g. Juravenator, Sinornithosaurus), although reticulate scales were probably retained on the mani and pedes of many theropods with a heavy plumage. Feathers and filamentous structures appear to have replaced widespread scaly integuments in maniraptorans. Theropod skin, and that of dinosaurs more broadly, remains a virtually untapped area of study and the appropriation of commonly used techniques in other palaeontological fields to the study of skin holds great promise for future insights into the biology, taphonomy and relationships of these extinct animals.

Hendrickx, C., Mateus O., & Buffetaut E. (2016).  Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.. PLoS ONE. 11, e0144695., 01, Number 1: Public Library of Science Abstracthendrickx_et_al_2016_morphofunctional_analysis_of_the_quadrate_of_spinosauridae_dinosauria.pdfWebsite

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.

Hendrickx, C., Mateus O., & Buffetaut E. (2016).  Morphofunctional Analysis of the Quadrate of Spinosauridae (Dinosauria: Theropoda) and the Presence of Spinosaurus and a Second Spinosaurine Taxon in the Cenomanian of North Africa.. PLoS ONE. 11, e0144695., 01, Number 1: Public Library of Science AbstractWebsite

Six quadrate bones, of which two almost certainly come from the Kem Kem beds (Cenomanian, Upper Cretaceous) of south-eastern Morocco, are determined to be from juvenile and adult individuals of Spinosaurinae based on phylogenetic, geometric morphometric, and phylogenetic morphometric analyses. Their morphology indicates two morphotypes evidencing the presence of two spinosaurine taxa ascribed to Spinosaurus aegyptiacus and? Sigilmassasaurus brevicollis in the Cenomanian of North Africa, casting doubt on the accuracy of some recent skeletal reconstructions which may be based on elements from several distinct species. Morphofunctional analysis of the mandibular articulation of the quadrate has shown that the jaw mechanics was peculiar in Spinosauridae. In mature spinosaurids, the posterior parts of the two mandibular rami displaced laterally when the jaw was depressed due to a lateromedially oriented intercondylar sulcus of the quadrate. Such lateral movement of the mandibular ramus was possible due to a movable mandibular symphysis in spinosaurids, allowing the pharynx to be widened. Similar jaw mechanics also occur in some pterosaurs and living pelecanids which are both adapted to capture and swallow large prey items. Spinosauridae, which were engaged, at least partially, in a piscivorous lifestyle, were able to consume large fish and may have occasionally fed on other prey such as pterosaurs and juvenile dinosaurs.

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Rotatori, F. M., Quaranta M., Bertozzo F., Hübner T., Camilo B., Mateus O., & Moreno-Azanza M. (2023).  Hadrosaur-like vascularisation in the dentary of an early diverging iguanodontian dinosaur. Historical Biology. 1-6.: Taylor & Francis AbstractWebsite

ABSTRACTVirtual palaeontology is a growing field, leading palaeontologists to better understand the microanatomy of many extinct species. The application of techniques such as CT and μCT-scanning allows the researchers to study micro-anatomical features in a non-invasive way and make inferences on the palaeobiology of animals. Dinosaurs have been extensively studied using these techniques, with particular focus on the microanatomy of the cranium, whereas relatively little is known of other cranial elements, such as the lower jaw. Here, we aim to fill this gap, describing the microanatomy of the specimen ML 768, an isolated dentary belonging to a dryosaurid iguanodontian dinosaur from the Upper Jurassic of Lourinhã Fm. The dentary ML 768 was subjected to μCT-scanning, and subsequently the data were segmented in Avizo and rendered in Blender. We identified functional and replacement teeth, recognising remnants of old replacement cycles. Furthermore, we mapped a rich neurovascular network present in the dentary and compared it with reference literature. We found that the high vascularisation is shared with other cerapodan dinosaurs with high tooth replacement rates, although homoeostasis may have also played a role in the development of this condition. Further evidence is needed to appreciate the macroevolutionary significance of these findings.

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Hayashi, S., Redelstorff R., Mateus O., Watabe M., & Carpenter K. (2014).  Gigantism of stegosaurian osteoderms. Journal of Vertebrate Paleontology. Program and Abstracts, 2014, 145.hayashi_et_al_2014_gigantism_of_stegosaurian_osteoderms.pdf
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Mateus, O., Callapez P. M., Polcyn M. J., Schulp A. S., Gonçalves A. O., & Jacobs L. L. (2019).  The Fossil Record of Biodiversity in Angola Through Time: A Paleontological Perspective. (Huntley, Brian J., Russo, Vladimir, Lages, Fernanda, Ferrand, Nuno, Ed.).Biodiversity of Angola: Science & Conservation: A Modern Synthesis. 53–76.: Springer International Publishing Abstractmateus2019_chapter_thefossilrecordofbiodiversityi.pdf

This chapter provides an overview of the alpha paleobiodiversity of Angola based on the available fossil record that is limited to the sedimentary rocks, ranging in age from Precambrian to the present. The geological period with the highest paleobiodiversity in the Angolan fossil record is the Cretaceous, with more than 80{%} of the total known fossil taxa, especially marine molluscs, including ammonites as a majority among them. The vertebrates represent about 15{%} of the known fauna and about one tenth of them are species firstly described based on specimens from Angola.

Mateus, O., Callapez P. M., Polcyn M. J., Schulp A. S., Gonçalves A. O., & Jacobs L. L. (2019).  The Fossil Record of Biodiversity in Angola Through Time: A Paleontological Perspective. (Huntley, Brian J., Russo, Vladimir, Lages, Fernanda, Ferrand, Nuno, Ed.).Biodiversity of Angola: Science {&} Conservation: A Modern Synthesis. 53–76., Cham: Springer International Publishing Abstract

This chapter provides an overview of the alpha paleobiodiversity of Angola based on the available fossil record that is limited to the sedimentary rocks, ranging in age from Precambrian to the present. The geological period with the highest paleobiodiversity in the Angolan fossil record is the Cretaceous, with more than 80{%} of the total known fossil taxa, especially marine molluscs, including ammonites as a majority among them. The vertebrates represent about 15{%} of the known fauna and about one tenth of them are species firstly described based on specimens from Angola.

Mallison, H., Schwarz-Wings D., Tsai H., Holliday C., & Mateus O. (2014).  Fossil longbone cartilage preserved in stegosaurs?. Journal of Vertebrate Paleontology. Program and Abstracts, 2014, 176.mallison_et_al._2014_fossil_longbone_cartilage_preserved_in_stegosaurs.pdf
Araújo, R., Castanhinha R., Martins R. M. S., Mateus O., Hendrickx C., Beckmann F., Schell N., & Alves L. C. (2013).  Filling the gaps of dinosaur eggshell phylogeny: Late Jurassic Theropod clutch with embryos from Portugal. Scientific Reports. 3(1924), , 2013/05/30/onlin: Macmillan Publishers Limited. All rights reserved Abstractaraujo_et_al_2013_filling_the_gaps_of_dinosaur_eggshell_phylogeny_late_jurassic_theropod_clutch_with_embryos_from_portugal.pdf

The non-avian saurischians that have associated eggshells and embryos are represented only by the sauropodomorph Massospondylus and Coelurosauria (derived theropods), thus missing the basal theropod representatives. We report a dinosaur clutch containing several crushed eggs and embryonic material ascribed to the megalosaurid theropod Torvosaurus. It represents the first associated eggshells and embryos of megalosauroids, thus filling an important phylogenetic gap between two distantly related groups of saurischians. These fossils represent the only unequivocal basal theropod embryos found to date. The assemblage was found in early Tithonian fluvial overbank deposits of the Lourinhã Formation in West Portugal. The morphological, microstructural and chemical characterization results of the eggshell fragments indicate very mild diagenesis. Furthermore, these fossils allow unambiguous association of basal theropod osteology with a specific and unique new eggshell morphology.

Araújo, R., Castanhinha R., Martins R. M. S., Mateus O., Hendrickx C., Beckmann F., Schell N., & Alves L. C. (2013).  Filling the gaps of dinosaur eggshell phylogeny: Late Jurassic Theropod clutch with embryos from Portugal. Scientific Reports. 3, , may: Nature Publishing Group AbstractWebsite
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Hendrickx, C., Mateus O., Araújo R., & Choiniere J. (2019).  The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica. 22(3), 1-110. Abstractthe_distribution_of_dental_features_in_non-avian_t.pdfWebsite

Isolated theropod teeth are some of the most common fossils in the dinosaur fossil record and are continually reported in the literature. Recently developed quantitative methods have improved our ability to test the affinities of isolated teeth in a repeatable framework. But in most studies, teeth are diagnosed on qualitative characters. This can be problematic because the distribution of theropod dental characters is still poorly documented, and often restricted to one lineage. To help in the identification of isolated theropod teeth, and to more rigorously evaluate their taxonomic and phylogenetic potential, we evaluated dental features in two ways. We first analyzed the distribution of 34 qualitative dental characters in a broad sample of taxa. Functional properties for each dental feature were included to assess how functional similarity generates homoplasy. We then compiled a quantitative data matrix of 145 dental characters for 97 saurischian taxa. The latter was used to assess the degree of homoplasy of qualitative dental characters, address longstanding questions on the taxonomic and biostratigraphic value of theropod teeth, and explore the major evolutionary trends in the theropod dentition.

In smaller phylogenetic datasets for Theropoda, dental characters exhibit higher levels of homoplasy than non-dental characters, yet they still provide useful grouping information and optimize as local synapomorphies of smaller clades. In broader phylogenetic datasets, the degree of homoplasy displayed by dental and non-dental characters is not significantly different. Dental features on crown ornamentations, enamel texture and tooth microstructure have significantly less homoplasy than other dental features and can be used to identify many theropod taxa to ‘family’ or ‘sub-family’ level, and some taxa to genus or species. These features should, therefore, be a priority for investigations seeking to classify isolated teeth.

Our observations improve the taxonomic utility of theropod teeth and in some cases can help make isolated teeth useful as biostratigraphic markers. This proposed list of dental features in theropods should, therefore, facilitate future studies on the systematic paleontology of isolated teeth.

Hendrickx, C., Mateus O., Araújo R., & Choiniere J. (2019).  The distribution of dental features in non-avian theropod dinosaurs: Taxonomic potential, degree of homoplasy, and major evolutionary trends. Palaeontologia Electronica. 22, , Number 3 Abstract
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Hendrickx, C., Mateus O., & Araújo R. (2014).  The dentition of Megalosauridae (Theropoda: Dinosauria). {APP}. : Polska Akademia Nauk Instytut Paleobiologii (Institute of Paleobiology, Polish Academy of Sciences) AbstractWebsite
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Hendrickx, C., Mateus O., & Araújo R. (2014).  The dentition of megalosaurid theropods, with a proposed terminology on theropod teeth. XII EAVP Meeting XII Annual Meeting of the European Association of Vertebrate Palaeontologists – Abstract Book. p. 75., Torino 24-28 June 2014hendrickx_et_al_2014_megalosaurid_teeth_eavp.pdf
Hendrickx, C., Mateus O., & Araújo R. (2015).  The dentition of megalosaurid theropods. Acta Palaeontologica Polonica. 60(3), 627–642. Abstracthendrickx_et_al_2015_theropod_teeth_app.pdfWebsite

Theropod teeth are particularly abundant in the fossil record and frequently reported in the literature. Yet, the dentition of many theropods has not been described comprehensively, omitting details on the denticle shape, crown ornamentation and enamel texture. This paucity of information has been particularly striking in basal clades, thus making identification of isolated teeth difficult, and taxonomic assignments uncertain. We here provide a detailed description of the dentition of Megalosauridae, and a comparison to and distinction from superficially similar teeth of all major theropod clades. Megalosaurid dinosaurs are characterized by a mesial carina facing mesiolabially in most mesial teeth, centrally positioned carinae on both most mesial and lateral crowns, a mesial carina terminating above the cervix, and short to well-developed interdenticular sulci between distal denticles. A discriminant analysis performed on a dataset of numerical data collected on the teeth of 62 theropod taxa reveals that megalosaurid teeth are hardly distinguishable from other theropod clades with ziphodont dentition. This study highlights the importance of detailing anatomical descriptions and providing additional morphometric data on teeth with the purpose of helping to identify isolated theropod teeth in the future.

Hayashi, S., Carpenter K., Watabe M., Mateus O., & Barsbold R. (2008).  Defensive weapons of thyreophoran dinosaurs: histological comparisons and structural differences in spikes and clubs of ankylosaurs and stegosaurs. 28 (3, Supplement), 89A-90A. Journal of Vertebrate Paleontology. 28, 89-90., Number Suppl. to 3 Abstract
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Hayashi, S., Carpenter K., Watabe M., Mateus O., & Barsbold R. (2008).  Defensive weapons of thyreophoran dinosaurs: histological comparisons and structural differences in spikes and clubs of ankylosaurs and stegosaurs. 28 (3, Supplement), 89A-90A. Journal of Vertebrate Paleontology. 28, 89–90., Number Suppl. to Abstract
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Hayashi, S., Carpenter K., Watabe M., Mateus O., & Barsbold R. (2008).  Defensive weapons of thyreophoran dinosaurs: histological comparisons and structural differences in spikes and clubs of ankylosaurs and stegosaurs. Journal of Vertebrate Paleontology. 28(3, Supplement), 89A-90A., Number Suppl. to 3 Abstracthayashi_et_al_2008_histology_stegosaurs_defensive_weapons_of_thyreophoran_dinosaurs-_histological_comparisons_and_structural_differences_in_spikes_and_clubs_of_ankylosaurs_and_stegosaurs.pdfWebsite

Thyreophoran dinosaurs have spike- and club-shaped osteoderms probably used for defensive weapons. The structural and histological variations have been little known. Here, we provide the comparisons of the internal structures in defensive weapons of ankylosaurs and stegosaurs, using spikes of a polacanthid (Gastonia) and a nodosaurid (Edmontonia), clubs of ankylosaurids (Saichania and Ankylosauridae indet. from Canada), and spikes of stegosaurids (Stegosaurus and Dacentrurus), which sheds light on understandings of evolutionary history and functional implications of defensive weapons in thyreophorans. In ankylosaurs, the structural and histological features of spikes and clubs are similar with those of small osteoderms in having thin compact bones, thick cancellous bones with large vascular canals, and abundant collagen fibers. A previous study demonstrated that each of three groups of ankylosaurs (polacanthid, nodosaurid, and ankylosaurid) has distinct characteristic arrangements of collagen fibers in small osteoderms. This study shows that spikes and clubs of ankylosaurs maintain the same characteristic features for each group despite of the differences in shapes and sizes. Conversely, the spike-shaped osteoderms in primitive (Dacentrurus) and derived (Stegosaurus) stegosaurids have similar structure to each other and are significantly different from the other types of stegosaur osteoderms (throat bony ossicles and plates) in having thick compact bones with a medullary cavity. These lack abundant collagen fibers unlike ankylosaur osteoderms. The spikes of ankylosaurs and stegosaurs are similar in shape, but their structural and histological features are different in having unique structures of collagen fibers for ankylosaurs and thick compact bones for stegosaurs, providing enough strength to have large spikes and to use them as defensive weapons. Although the shapes of ankylosaur clubs are different from spikes, the internal structures are similar, suggesting that ankylosaurs maintain similar structures despite of different shapes in osteoderms. These results indicate that ankylosaurs and stegosaurs used different strategies independently to evolve defensive weapons.

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Hansen, B. B., Milàn J., Clemmensen L. B., Adolfssen J. S., Estrup E. J., Klein N., Mateus O., & Wings O. (2016).  Coprolites from the Late Triassic Kap Stewart Formation, Jameson Land, East Greenland: morphology, classification and prey inclusions. Geological Society, London, Special Publications. 434(1), 49-69. Abstracthansen_et_al_2015_coprolites_from_the_late_triassic_kap_stewart_formation_jameson_land_east_greenland.pdfWebsite

A large collection of vertebrate coprolites from black lacustrine shales in the Late Triassic (Rhaetian–Sinemurian) Kap Stewart Formation, East Greenland is examined with regard to internal and external morphology, prey inclusions, and possible relationships to the contemporary vertebrate fauna. A number of the coprolites were mineralogically examined by X-ray diffraction (XRD), showing the primary mineral composition to be apatite, clay minerals, carbonates and, occasionally, quartz in the form of secondary mineral grains. The coprolite assemblage shows multiple sizes and morphotypes of coprolites, and different types of prey inclusions, demonstrating that the coprolite assemblage originates from a variety of different producers.Supplementary material: A description of the size, shape, structure, texture, contents and preservation of the 328 specimens is available at https://doi.org/10.6084/m9.figshare.c.2134335

Hansen, B. B., Milàn J., Clemmensen L. B., Adolfssen J. S., Estrup E. J., Klein N., Mateus O., & Wings O. (2015).  Coprolites from the Late Triassic Kap Stewart Formation, Jameson Land, East Greenland: morphology, classification and prey inclusions. Geological Society, London, Special Publications. 434, AbstractWebsite

A large collection of vertebrate coprolites from black lacustrine shales in the Late Triassic (Rhaetian–Sinemurian) Kap Stewart Formation, East Greenland is examined with regard to internal and external morphology, prey inclusions, and possible relationships to the contemporary vertebrate fauna. A number of the coprolites were mineralogically examined by X-ray diffraction (XRD), showing the primary mineral composition to be apatite, clay minerals, carbonates and, occasionally, quartz in the form of secondary mineral grains. The coprolite assemblage shows multiple sizes and morphotypes of coprolites, and different types of prey inclusions, demonstrating that the coprolite assemblage originates from a variety of different producers.Supplementary material: A description of the size, shape, structure, texture, contents and preservation of the 328 specimens is available at https://doi.org/10.6084/m9.figshare.c.2134335

Tschopp, E., Brinkman D., Henderson J., Turner M. A., & Mateus O. (2018).  Considerations on the replacement of a type species in the case of the sauropod dinosaur Diplodocus Marsh, 1878. Geology of the Intermountain West. 5, 245-262.tschoppetal2018.pdf
Tschopp, E., Brinkman D., Henderson J., Turner M. A., & Mateus O. (2018).  Considerations on the replacement of a type species in the case of the sauropod dinosaur Diplodocus Marsh, 1878. Geology of the Intermountain West. 5, 245-262. Abstract
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Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ruiz-Omeñaca J. I., Havlik P., Lepage Y., & de Andrade M. B. (2015).  Addendum to ‘Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia)’. Royal Society Open Science. 2, , Number 2: The Royal Society Abstractyoung_et_al_2015_addendum_to_revision_of_the_late_jurassic_teleosaurid_genus_machimosaurus_crocodylomorpha_thalattosuchia.pdfWebsite

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Young, M. T., Hua S., Steel L., Foffa D., Brusatte S. L., Thüring S., Mateus O., Ruiz-Omeñaca J. I., Havlik P., Lepage Y., & De Andrade M. B. (2015).  Addendum to ?Revision of the Late Jurassic teleosaurid genus Machimosaurus (Crocodylomorpha, Thalattosuchia)?. Royal Society Open Science. 2, , Number 2 Abstract
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Hendrickx, C., & Mateus O. (2014).  Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa. 3759, 1-74. Abstracthendrickx__mateus_2014._abelisauridae_dinosauria_theropoda_from_the_late_jurassic_of_portugal.pdf

Theropod dinosaurs form a highly diversified clade, and their teeth are some of the most common components of the Mesozoic dinosaur fossil record. This is the case in the Lourinhã Formation (Late Jurassic, Kimmeridgian-Tithonian) of Portugal, where theropod teeth are particularly abundant and diverse. Four isolated theropod teeth are here described and identified based on morphometric and anatomical data. They are included in a cladistic analysis performed on a data matrix of 141 dentition-based characters coded in 60 taxa, as well as a supermatrix combining our dataset with six recent datamatrices based on the whole theropod skeleton. The consensus tree resulting from the dentition-based data matrix reveals that theropod teeth provide reliable data for identification at approximately family level. Therefore, phylogenetic methods will help identifying theropod teeth with more confidence in the future. Although dental characters do not reliably indicate relationships among higher clades of theropods, they demonstrate interesting patterns of homoplasy suggesting dietary convergence in (1) alvarezsauroids, therizinosaurs and troodontids; (2) coelophysoids and spinosaurids; (3) compsognathids and dromaeosaurids; and (4) ceratosaurids, allosauroids and megalosaurids.

Based on morphometric and cladistic analyses, the biggest tooth from Lourinhã is referred to a mesial crown of the megalosaurid Torvosaurus tanneri, due to the elliptical cross section of the crown base, the large size and elongation of the crown, medially positioned mesial and distal carinae, and the coarse denticles. The smallest tooth is identified as Richardoestesia, and as a close relative of R. gilmorei based on the weak constriction between crown and root, the “eight-shaped” outline of the base crown and, on the distal carina, the average of ten symmetrically rounded denticles per mm, as well as a subequal number of denticles basally and at mid-crown. Finally, the two medium-sized teeth belong to the same taxon and exhibit pronounced interdenticular sulci between distal denticles, hooked distal denticles for one of them, an irregular enamel texture, and a straight distal margin, a combination of features only observed in abelisaurids. They provide the first record of Abelisauridae in the Jurassic of Laurasia and one of the oldest records of this clade in the world, suggesting a possible radiation of Abelisauridae in Europe well before the Upper Cretaceous.